1. Hearing

2. Outer ear
The ear has three functional parts. The main part of the external ear, the auricle, captures sound efficiently and sends it into the ear canal. Our capacity to localize sounds in space, especially along the vertical axis, depends critically upon the sound - gathering properties of the external ear.

3. Middle ear
Middle ear transmits mechanical energy to the ear's receptive organ. The three tiny ossicles, or bones: the malleus, or hammer; the incus, or anvil; and the stapes, or stirrup pass the motions of the tympanic membrane (eardrum) to the oval window – the bony covering of the cochlea.

4. Reflection and transmission at the eardrum
environment (air)
inner ear (water)
Amplitude reflection and transmission coefficients:
Energy reflection and transmission coefficients:
Z – mechanical impedance (= rv)
r - density
v – wave propagation speed

5. Reflection and transmission at the eardrum
rair = 1.21 kg/m3
vair = 340 m/s
rwater = 103 kg/m3
vwater = 1480 m/s
This values yield energy reflection and transmission coefficients:
T = 1 – R =
R =
0.1% energy transmitted
~99.9% is reflected
If only 1 part in 1000 makes it through, the loss is:
10log101/1000 = -30 dB

6. The middle ear
Purpose of the bones in the middle ear is to serve as amplifying system for the transmission of the eardrum vibrations towards the inner ear.
Tympanic membrane
Malleus
Incus
Stapes
~ 55 mm2
~ 3.2 mm2
Area ratio about 17:1 gives 17-fold sound pressure increase. It represents an increase in decibels of:
20log1017/1 = 25dB

7. Sound pressure level increase is approximately: 20log109/7 = 2 dB
The middle ear
Incus
7 mm
Malleus
9 mm
Malleus/Incus length ratio is 9:7 so force and hence the pressure is increased by 1.3:1
Sound pressure level increase is approximately:
20log109/7 = 2 dB

8. Eardrum displacement at threshold of hearing
Acoustic travelling wave
Total wave energy
(Epot = Ekin)
Wave intensity
Wave intensity at the threshold of hearing 0 dB
Wave amplitude
Wave amplitude at 440 Hz
Hydrogen atom diameter
The eardrum displacement at threshold of hearing is ~ 1/10 the diameter of a hydrogen atom!

9. The threshold of hearing
At a threshold of 3000 Hz (the frequency to which we are most sensitive – important for speech) the eardrum displacement is about one tenth of the diameter of a single hydrogen atom.
In a very quiet setting, blood can actually be heard flowing through the vessels near the ear. If ears were much more sensitive, we would be hearing noise generated by air molecules colliding with the eardrum.
A threshold vibration deflects auditory receptors in an inner ear (stereocilia) through an angle of only about degree. Bending the Empire State Building through such an angle would deflect its top by less than 2.5 cm.
From this threshold we can hear over a 10 million–fold range of sound pressure levels before sounds become painfully loud. Because it is cumbersome to keep track of all the zeros in such large numbers, a logarithmic decibel scale was introduced.

10. Sound Intensity and Sound Pressure Levels
Human ear responds to sound intensities in the range of 10-12– 100 W/m2. Sound intensity level is defined as:
where:
– sound intensity level
I – sound intensity
I0 – standard reference sound intensity 10–12 W/m2
In such scale 10-fold increase in sound intensity, gives 10 dB increase in sound volume (intensity level).
Sound pressure level (SPL) or sound level is a logarithmic measure of the effective sound pressure of a sound relative to a reference value:

11. Sound Intensity and Sound Pressure Levels
Intensity dB Pressure Examples
108 W/m *105 Pa Volcano erruption
102 W/m *102 Pa Jet aircraft, 50 m
1 W/m *101 Pa Rock concert
10-2 W/m Pa Disco, 1m from the speaker
10-4 W/m *10-1 Pa Highway, 5 m
10-6 W/m *10-2 Pa Speech, 1 m
10-8 W/m *10-3 Pa Background in quiet library
10-10 W/m *10-4 Pa Background in TV studio
10-12 W/m *10-5 Pa Hearing threshold
Threshold of pain 130 dB
Discomfort threshold, hearing damage possible 120 dB
limit Apple iPod volume in Europe 85 dB

12. Building a cochlea

13. Building a cochlea

14. Building a cochlea

15. Building a cochlea

16. Building a cochlea

17. How cochlea works ? High frequency Low frequency
Cochlea in the inner ear is shaped like a snail shell. Net effect is to boost pressure created by sound so that the inner ear fluid can be displaced. Its spiral shape effectively boosts the strength of the vibrations caused by sound, especially for low frequencies.

18. Cochlea and basilar membrane
Cochlea contains basilar membrane, which separates two liquid-filled tubes that run along the cochlea, the scala media and the scala tympani. Basilar membrane is narrow at the base and widens at the tip. Different frequencies are coded by the position along the membrane – high frequencies displace the membrane at the base, low frequencies displace the membrane at the apex.

19. Helmholtz’s resonance theory of hearing
Different frequencies of sound are encoded by their precise position along the basilar membrane. Short strings at the base would resonate in response to high notes, and the long strings (at the apex) to low notes.

20. The travelling wave theory - Von Bekesy (1928). Nobel 1961
Georg von Békésy (1899 – 1972)
A. The sound pressure applied to the oval window produces travelling wave along the basilar membrane. C. The peak displacements for high frequencies are toward the base, and for low frequencies are toward the apex.

21. Evidence for active amplification mechanism
Theory of van Beksy contradicts our daily experience. Envelopes of the travelling waves are wide while we are able to hear pure tones. There must be some additional mechanism for tunning of the auditory system to the sound frequency. The frequency response of individual cochlear nerve fibers depends on a combination of the mechanical properties of the basilar membrane and an active amplification mechanism.
Effect of cochlear amplifier. (c) The peak due to cochlear amplifier.(d) Amplitude of the passive movement of basilar membrane with metabolic block applied of the cochlear amplifier.

22. The Organ of Corti
The organ of Corti is the receptor organ of the inner ear, which actively amplifies the travelling wave. It contains the hair cells and a variety of supporting cells.
A. Light micrograph of a human organ of Corti. B. Drawing of a cross section of the organ of Corti.

23. Two types of hair cells
Scanning electron micrographs of the organ of Corti after removal of the tectorial membrane. Inner hair cells are arranged in the single row. Outer hair cells are arranged in the three rows and the stereocilia of each cell are arranged in a V configuration.

24. Inner hair cells are sensory cells; outer hair cells are amplifiers
95 % of the fibers of the auditory nerve arise from the inner hair cells. It suggests that the inner hair cells are actual sensory receptors.
Innervation pattern: nerve fibers connect to the 3500 IHC, while 1000 nerve fibers connect to the OHC. The IHC are the main sites of auditory transduction.

25. Outer hair cells
Outer hair cell may change its length in response to stimulus. The ‘motor’ is a transmembrane protein that mechanically contracts and elongates the cell. The molecule, discovered in 2000 is called ‘prestin’.

26. Rock Around the Clock Hair Cell
An outer hair cell is being stimulated electrically by a patch pipette which enters from the lower left. The cell’s potential is changed by by plugging Walkman into the input socket of the electrophysiology amplifier.
(from:

27. The cochlear amplifier
Shape changes of the outer hair cells due to rapidly oscillating membranne potential contribute to movement of the tectorial and basilar membranes. Inner hair cells are stimulated by the relative movements between these membranes. It is presumed that this mechanism contributes to the active tunning of hair cells responses.

28. Otoacoustic emission
An otoacoustic emission (OAE) is a sound which is generated from within the inner ear. There are two types of otoacoustic emissions: spontaneous otoacoustic emissions, which can occur without external stimulation, and evoked otoacoustic emissions, which require an evoking stimulus. Most probably, otoacoustic emissions are produced by the cochlear outer hair cells as they expand and contract. Otoacoustic emissions are the basis of a simple, non-invasive, test for hearing defects in newborn babies.
An example of multifrequency spontaneous otoacoustic emissions recorded from a 48-year-old woman with normal hearing. The black spikes represent the response above the noise floor.
An example of evoked otoacoustic emissions and their spectra. Evoked otoacoustical emissions are evidence for a cochlear amplifier.

29. Mechanism of frequency tunning 2 – dependence on the location
Many properties of IHC and OHC vary with the position along the cochlea. These differences are likely to be correlated with the differing frequencies that are processed along the cochlea, but the significance of these changes is still not understood.

30. Efferent fibers
In addition to afferent fibers, the auditory nerve also contains efferent fibers, which arises from cells in the brain-stem. Efferent fibers inhibit mainly outer hair cells by hyperpolarizing the hair cells membrane. It reduces the motor output of the outer hair cells and reduces the movement of the tectorial and basilar membranes and the sensory response of the inner hair cells. Its role is assumed to be a protection against overstimulation.

31. Tunning curves
Tuning curves for cochlear hair cells. To construct a curve, the experimenter presents sound at each frequency at increasing amplitudes until the cell produces a criterion response, here 1 mV. The curve thus reflects the threshold of the cell for stimulation at a range of frequencies. Each cell is most sensitive to a specific frequency. The threshold rises briskly (sensitivity falls abruptly) as the stimulus frequency is raised or lowered.

32. Auditory pathways Auditory pathways: - Cochlea
Cochlear nuclei (brain-stem)
Superior olivary nuclei (brain-stem)
Inferior colliculus (brain-stem)
Medial geniculate nuclei (thalamus)
Auditory cortex
Left
Auditory
cortex
Right
Auditory
cortex
Medial geniculate nucleus
Cochlea
Inferior colliculus
Auditory
nerve fiber
Superior
Olivary
nucleus
Ipsilateral
Cochlear
nucleus

33. Types of cells in the cochlear nuclei
Auditory nerve fibers terminate in the cochlear nuclei (CN) on different types of cells with different response properties. Responses to a tone burst of 50 ms are shown. The ‘Primary-like’ preserve the envelope of the input signal, the ‘Pauser’ and the’Chopper’ provide for differentation between onset and ensuing phases of the tone, the ‘On’ cells signal the onset or timing of a sound. Each cell type represents an abstraction of one particular feature of the input. Different functional properties are processed and transmitted in parallel pathways. In humans, the receptor potentials of certain hair cells and the action potentials of their associated auditory nerve fiber can follow stimuli of up to about 3 kHz in a one-to-one fashion.

34. Sound localization in medial superior olive nuclei
Diagram illustrating how the MSO computes the location of a sound by interaural time differences. A given MSO neuron responds most strongly when the two inputs arrive simultaneously, as occurs when the contralateral and ipsilateral inputs precisely compensate (via their different lengths) for differences in the time of arrival of a sound at the two ears. The systematic (and inverse) variation in the delay lengths of the two inputs creates a map of sound location: In this model, E would be most sensitive to sounds located to the left, and A to sounds from the right; C would respond best to sounds coming from directly in front of the listener. Psychophysical experiments show that humans can actually detect interaural time differences as small as 10 microseconds; This sensitivity translates into an accuracy for sound localization of about 1°. Interaural time differences are used to localize the source for frequencies below 3 kHz .

35. Sound localization in lateral superior olive nuclei
Lateral superior olive neurons encode sound location through interaural intensity differences. LSO neurons receive direct excitation from the ipsilateral cochlear nucleus; input from the contralateral cochlear nucleus is relayed via inhibitory interneurons in the MNTB (medial nucleus of the trapezoid body). This excitatory/inhibitory interaction results in a net excitation of the LSO on the same side of the body as the sound source. In contrast, sounds arising from in front of the listener, will silence the LSO output. Interaural intensity differences are used to localize the source for frequencies above 2 kHz .

36. Tonotopic organisation
The basilar membrane in the cochlea is tonotopically organized. The tonotopic organization is retained at all levels of the central auditory system.

37. The auditory cortex
Diagram showing the brain in left lateral view, including the depths of the lateral sulcus, where part of the auditory cortex occupying the superior temporal gyrus normally lies hidden. The primary auditory cortex (A1) is shown in blue; the surrounding belt areas of the auditory cortex are in red. The primary auditory cortex has a tonotopic organization, as shown in the blowup diagram of a segment of A1. The Wernicke's area shown in green is a region important in comprehending speech. It is just posterior to the primary auditory cortex.

38. Tonotopic mapping in auditory cortex
Tonotopic mapping in human primary auditory cortex, demonstrated by applying 500-msec tone bursts at five different sound frequencies to one ear while mapping the magnetic field changes produced by neuronal current flow in primary auditory cortex. Location of the primary auditory cortex in the transverse temporal gyri is visible.

39. Tritone paradox frequency sequence
Shepard tones. Each square in the figure indicates a tone, with any set of squares in vertical alignment together making one Shepard tone. The color of each square indicates the loudness of the note, with purple being the quietest and green the loudest. Overlapping notes in one Shepard tone are exactly one octave apart. A pair of Shepard tones separated by an interval of a tritone (half an octave) produces the tritone paradox. They may be heard as either descending or ascending and constitute a bistable figure in auditory domain.